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There is a well-established discrepancy between paleontological and molecular data regarding the timing of the origin and diversification of placental mammals. Molecular estimates place interordinal diversification dates in the Cretaceous, while no unambiguous crown placental fossils have been found prior to the end-Cretaceous mass extinction. Here, the completeness of the eutherian fossil record through geological time is evaluated to assess the suggestion that a poor fossil record is largely responsible for the difference in estimates of placental origins. The completeness of fossil specimens was measured using the character completeness metric, which quantifies the completeness of fossil taxa as the percentage of phylogenetic characters available to be scored for any given taxon. Our data set comprised 33 published cladistic matrices representing 445 genera, of which 333 were coded at the species level.
Therewas no significant difference in eutherian completeness across the Cretaceous/Paleogene (K/Pg) boundary. This suggests that the lack of placental mammal fossils in the Cretaceous is not due to a poor fossil record but more likely represents a genuine absence of placental mammals in the Cretaceous. This result supports the “explosivemodel” of early placental evolution, whereby placentalmammals originated around the time of the K/Pg boundary and diversified soon after.
No correlation was found between the completeness pattern observed in this study and those of previous completeness studies on birds and sauropodomorph dinosaurs, suggesting that different factors affect the preservation of these groups. No correlations were found with various isotope proxy measures, but Akaike information criterion analysis found that eutherian character completeness metric scores were best explained by models involving the marine-carbonate strontium-isotope ratios (87Sr/86Sr), suggesting that tectonic activity might play a role in controlling the completeness of the eutherian fossil record.
Whether the evolutionary dynamics of one group of organisms influence that of another group of organisms over the vast timescale of the geological record is a difficult question to tackle. This is not least because multiple factors can influence or mask the effects of potential driving forces on evolutionary dynamics of the focal group. Here, we show how an approach amenable to causality inference for time series, linear stochastic differential equations (SDEs), can be used in a multivariate fashion to shed light on driving forces of diversification dynamics across the Phanerozoic. Using a new, enhanced stepwise search algorithm, we searched through hundreds of models to converge on a model that best describes the dynamic relationships that drove brachiopod and bivalve diversification rates. Using this multivariate framework, we characterized a slow process (half-life of c. 42 Myr) that drove brachiopod extinction. This slow process has yet to be identified from the geological record. Using our new framework for analyzing multiple linear SDEs, we also corroborate our previous findings that bivalve extinction drove brachiopod origination in the sense that brachiopods tended to diversify at a greater rate when bivalves were removed from the system. It is also very likely that bivalves “self-regulate” in the sense that bivalve extinctions also paved the way for higher bivalve origination rates. Multivariate linear SDEs as we presented them here are likely useful for studying other dynamic systems whose signatures are preserved in the paleontological record.
In this study we focused on the dynamics of encrusting assemblages preserved on brachiopod hosts collected from upper Frasnian and lower Famennian deposits of the Central Devonian Field, Russia. Because the encrusted brachiopods come from deposits bracketing the Frasnian/Famennian (F/F) boundary, the results also shed some light on ecological differences in encrusting communities before and after the Frasnian—Famennian (F-F) event. To explore the diversity dynamics of encrusting assemblages, we analyzed more than 1300 brachiopod valves (substrates) from two localities. Taxon accumulation plots and shareholder quorumsubsampling (SQS) routines indicated that a reasonably small sample of brachiopod host valves (n=50) is sufficient to capture themajority of the encrusting genera recorded at a given site. The richness of encrusters per substrate declined simultaneously with the number of encrusting taxa in the lower Famennian, accompanied by a decrease in epibiont abundance, with a comparable decrease in mean encrustation intensity (percentage of bioclasts encrusted by one or more epibionts). Epibiont abundance and occupancy roughlymirror each other. Strikingly, few ecological characteristics are correlated with substrate size, possibly reflecting random settlement of larvae. Evenness, which is negatively correlated with substrate size, shows greater within-stage variability among samples than between Frasnian and Famennian intervals and may indicate the instability of early Famennian biocenoses following the faunal turnover. The occurrence distribution of encrusters points to nonrandomassociations and exclusions among several encrusting taxa. However, abundance and occupancy of microconchids remained relatively stable throughout the sampled time interval. The notable decline in abundance (∼60%) and relatively minor decline in diversity (∼30%) suggest jointly that encrusting communities experienced ecological collapse rather than a major mass extinction event. The differences between the upper Frasnian and lower Famennian encrusting assemblages may thus record a turnover associated with the F-F event.
Understanding the drivers of macroevolutionary trends through the Phanerozoic has been a central question in paleobiology. Increasingly important is understanding the regional and environmental variation of macroevolutionary patterns and how they are reflected at the global scale. Here we test the role of biotic interactions on regional ecological patterns during the Mesozoic marine revolution. We test for escalatory trends in Jurassic marine benthic macroinvertebrate ecosystems using occurrence data from the Paleobiology Database parsed by region and environment. The escalation hypothesis posits that taxonomic groups that could adapt to intense predation and bioturbation proliferated, whereas groups unable to adapt were reduced in diversity and abundance or driven to extinction. We tested this hypothesis in five regions during Jurassic stages and among four depositional environments in Europe. Few escalatory trends were detected, although at least one escalatory trend was observed in every region, with the greatest number and strongest trends observed in Europe. These trends include increases in shallow infauna and cementing epifauna and occurrences of facultatively mobile invertebrates and decreases in pedunculate, free-lying, and sessile epifauna. Within Europe, escalatory trends occur in shallow-water environments but also in deeper-water environments, where they are predicted not to occur. When regional trends are aggregated, trends in Europe drive the global signal. The results of this study suggest that while evidence of escalation is rare globally, it is plausible that escalation drove macroevolutionary patterns in Europe. Furthermore, these results underline the need to dissect global fossil data at the regional scale to understand global macroevolutionary dynamics.
Two end-member models are proposed to explain marine biotic responses to greenhouse conditions. Global warming and increasing sea level may: (1) promote dispersal of marine species, leading to larger geographic ranges and decreased speciation and biodiversity; or (2) result in formation of isolated epicontinental basins that host endemic radiations, leading to smaller geographic ranges and increased speciation and biodiversity. The Cenomanian—Turonian (C-T) interval, marked by greenhouse warming, sea-level rise, ocean anoxia, and biotic turnover, presents an opportunity to test these two end-member models. In particular, how cephalopods responded to these global changes has not been clear. A global database of 7262 cephalopod occurrences was used to evaluate biodiversity changes through the C-T interval. Both species- and genus-level diversity peaked in the late Cenomanian. The global diversity drop across the C/T boundary was modest; rather, diversity was low during the middle Cenomanian and middle Turonian, times of brief cooling. Regional variations in diversity responses may reflect the degree and timing of environmental perturbations within different oceanographic settings. Surprisingly, cephalopod faunas in the European Platform, Western Interior, and South Atlantic all shifted equatorward across the C/T boundary, whereas other regions saw no change in latitudinal distributions. Global generic geographic ranges did not change through the C-T interval, but the percentage of cosmopolitan genera did increase significantly across the C/T, both globally and within the Western Interior and Europe, whereas cosmopolitans dropped in the Pacific and South Atlantic. Neither end-member model for biodiversity change in a greenhouse world is supported for C-T cephalopods, as diversity increased without an associated increase in geographic range. It may be that sea-level rise and global warming led to both endemic radiations in epicontinental basins and an increase in cosmopolitan taxa in some regions, demonstrating the importance of combining global and regional-scale analyses.
Sexual dimorphism is common in many extant animals, but it is difficult to demonstrate in fossil species. Working with material from the Late Cretaceous of the U.S. Coastal Plain, we herein analyze sexual dimorphism in ostracodes from the superfamily Cytheroidea, a group whose extant members have males that are relatively more elongate than females. We digitized outlines of more than 6000 individual ostracode valves or carapaces, extracted size (area) and shape (length-to-height ratio) information, and used finite mixture models to assess hypotheses of sexual dimorphism. Male and female clusters can be discerned in nearly all populations with sufficient data, resulting in estimates of size and shape dimorphismfor 142 populations across 106 species; an additional nine samples are interpreted to consist only of females. Dimorphism patterns varied across taxa, especially for body size: males range from 30% larger to 20% smaller than females. Magnitudes of sexual dimorphism are generally stable within species across time and space; we can demonstrate substantial evolutionary changes in dimorphism in only one species, Haplocytheridea renfroensis. Several lines of evidence indicate that patterns of sexual dimorphism in these ostracodes reflect male investment in reproduction, suggesting that this study system has the potential to capture variation in sexual selection through the fossil record.
The late Pleistocene megafaunal extinctions may have been the first extinctions directly related to human activity, but in North America the close temporal proximity of human arrival and the Younger Dryas climate event has hindered efforts to identify the ultimate extinction cause. Previous work evaluating the roles of climate change and human activity in the North American megafaunal extinction has been stymied by a reliance on geographic binning, yielding contradictory results among researchers. We used a fine-scale geospatial approach in combination with 95 megafaunal last-appearance and 75 human first-appearance radiocarbon dates to evaluate the North American megafaunal extinction. We used kriging to create interpolated first- and last-appearance surfaces from calibrated radiocarbon dates in combination with their geographic autocorrelation. We found substantial evidence for overlap between megafaunal and human populations in many but not all areas, in some cases exceeding 3000 years of predicted overlap. We also found that overlap was highly regional: megafauna had last appearances in Alaska before humans first appeared, but did not have last appearances in the Great Lakes region until several thousand years after the first recorded human appearances. Overlap in the Great Lakes region exceeds uncertainty in radiocarbon measurements ormethodological uncertainty and would be even greater with sampling-derived confidence intervals. The kriged maps of last megafaunal occurrence are consistent with climate as a primary driver in some areas, but we cannot eliminate human influence from all regions. The late Pleistocene megafaunal extinction was highly variable in timing and duration of human overlap across the continent, and future analyses should take these regional trends into account.
Patterns of turnover in the mammalian fossil record have been interpreted as reflecting “pulses” of originations and extinctions hypothesized to be driven by climate change. However, criteria for determining what constitutes a meaningful pulse have been idiosyncratic, and investigations of turnover patterns in mammals have yielded mixed results.
This study presents simple simulations of fossil records in which origination and extinction probabilities for each lineage are held constant. Nonetheless, the total number of turnover events per time bin varies stochastically, producing statistical “noise.” Various simulation and analytical assumptions are examined to determine their impact on the type I error rate (i.e., how often “pulses” are detected in a purely stochastic process).
Results suggest that simple analytical parameters (length of time bins and turnover-pulse criterion) have predictable and straightforward effects on false-positive rates. Furthermore, “pulses” of turnover of a magnitude similar to that observed in the terrestrial mammalian fossil record may be quite common under realistic analytical conditions.
The null turnover model offers a practical way to evaluate the significance of observed turnover events in future empirical studies of the fossil record. In evaluating the significance of a “pulse” of fossil origination or extinction events, analytical parameters can be explored using this null model to determine the approximate type I error rate for a set of parameters. Because false-positive rates are shown to be quite high, functional trait-based approaches may offer more reliable indicators of the impact of climate change on turnover dynamics.
Estimating biodiversity and its variations through geologic time is a notoriously difficult task, due to several taphonomic and methodological effects that make the reconstructed signal potentially distinct from the unknown, original one. Through a simulation approach, we examine the effect of a major, surprisingly still understudied, source of potential disturbance: the effect of time discretization through biochronological construction, which generates spurious coexistences of taxa within discrete time intervals (i.e., biozones), and thus potentially makes continuous- and discrete-time biodiversity curves very different. Focusing on the taxonomic-richness dimension of biodiversity (including estimates of origination and extinction rates), our approach relies on generation of random continuous-time richness curves, which are then time-discretized to estimate the noise generated by this manipulation. A broad spectrum of data-set parameters (including average taxon longevity and biozone duration, total number of taxa, and simulated time interval) is evaluated through sensitivity analysis. We show that the deteriorating effect of time discretization on the richness signal depends highly on such parameters, most particularly on average biozone duration and taxonomic longevity because of their direct relationship with the number of false coexistences generated by time discretization. With several worst-case but realistic parameter combinations (e.g., when relatively short-lived taxa are analyzed in a long-ranging biozone framework), the original and time-discretized richness curves can ultimately show a very weak to zero correlation, making these two time series independent. Based on these simulation results, we propose a simple algorithm allowing the back-transformation of a discrete-time taxonomic-richness data set, as customarily constructed by paleontologists, into a continuous-time data set. We show that the reconstructed richness curve obtained this way fits the original signal much more closely, even when the parameter combination of the original data set is particularly adverse to an effective time-discretized reconstruction.
Despite more than a century of interest, body-mass estimation in the fossil record remains contentious, particularly when estimating the body mass of taxa outside the size scope of living animals. One estimation approach uses humeral and femoral (stylopodial) circumferences collected from extant (living) terrestrial vertebrates to infer the body masses of extinct tetrapods through scaling models. When applied to very large extinct taxa, extant-based scaling approaches incur obvious methodological extrapolations leading some to suggest that they may overestimate the body masses of large terrestrial vertebrates. Here, I test the implicit assumption of such assertions: that a quadratic model provides a better fit to the combined humeral and femoral circumferences-to-body mass relationship. I then examine the extrapolation potential of these models through a series of subsetting exercises in which lower body-mass sets are used to estimate larger sets. Model fitting recovered greater support for the original linear model, and a nonsignificant second-degree term indicates that the quadratic relationship is statistically linear. Nevertheless, some statistical support was obtained for the quadratic model, and application of the quadratic model to a series of dinosaurs provides lower mass estimates at larger sizes that are more consistent with recent estimates using a minimum convex-hull (MCH) approach. Given this consistency, a quadratic model may be preferred at this time. Still, caution is advised; extrapolations of quadratic functions are unpredictable compared with linear functions. Further research testing the MCH approach (e.g., the use of a universal upscaling factor) may shed light on the linear versus quadratic nature of the relationship between the combined femoral and humeral circumferences and body mass.